(FIGURE 2.274. Skull of a fossil lemur (Adapis parisiensis,), from the Miocene at Quercy. A lateral view from the right, half natural size. B lower jaw, C lower molar, i incisors, c canines, p premolars, m molars.)
The manner in which these two kinds of vessels combine in the placenta, and the structure, form, and size of it, differ a good deal in the various Placentals; to some extent they give us valuable data for the natural classification, and therefore the phylogeny, of the whole of this sub-class. On the ground of these differences we divide it into two principal sections; the lower Placentals or Indecidua, and the higher Placentals or Deciduata.
To the Indecidua belong three important groups of mammals: the Lemurs (Prosimiae), the Ungulates (tapirs, horses, pigs, ruminants, etc.), and the Cetacea (dolphins and whales). In these Indecidua the villi are distributed over the whole surface of the chorion (or its greater part) either singly or in groups. They are only loosely connected with the mucous coat of the uterus, so that the whole foetal membrane with its villi can be easily withdrawn from the uterine depressions like a hand from a glove. There is no real coalescence of the two placentas at any part of the surface of contact. Hence at birth the foetal placenta alone comes away; the uterine placenta is not torn away with it.
The formation of the placenta is very different in the second and higher section of the Placentals, the Deciduata. Here again the whole surface of the chorion is thickly covered with the villi in the beginning. But they afterwards disappear from one part of the surface, and grow proportionately thicker on the other part. We thus get a differentiation between the smooth chorion (chorion laeve, Figure 2.273 chl) and the thickly-tufted chorion (chorion frondosum, Figure 2.273 chf). The former has only a few small villi or none at all; the latter is thickly covered with large and well-developed villi; this alone now constitutes the placenta. In the great majority of the Deciduata the placenta has the same shape as in man (Figures 1.197 and 1.200)—namely a thick, circular disk like a cake; so we find in the Insectivora, Chiroptera, Rodents, and Apes. This discoplacenta lies on one side of the chorion. But in the Sarcotheria (both the Carnivora and the seals, Pinnipedia) and in the elephant and several other Deciduates we find a zonoplacenta; in these the rich mass of villi runs like a girdle round the middle of the ellipsoid chorion, the two poles of it being free from them.
(FIGURE 2.275. The Slender Lori (Stenops gracilis) of Ceylon, a tail-less lemur.)
Still more characteristic of the Deciduates is the peculiar and very intimate connection between the chorion frondosum and the corresponding part of the mucous coat of the womb, which we must regard as a real coalescence of the two. The villi of the chorion push their branches into the blood-filled tissues of the coat of the uterus, and the vessels of each loop together so intimately that it is no longer possible to separate the foetal from the maternal placenta; they form henceforth a compact and apparently simple placenta. In consequence of this coalescence, a whole piece of the lining of the womb comes away at birth with the foetal membrane that is interlaced with it. This piece is called the "falling-away" membrane (decidua). It is also called the serous (spongy) membrane, because it is pierced like a sieve or sponge. All the higher Placentals that have this decidua are classed together as the "Deciduates." The tearing away of the decidua at birth naturally causes the mother to lose a quantity of blood, which does not happen in the Indecidua. The last part of the uterine coat has to be repaired by a new growth after birth in the Deciduates. (Cf. Figures 1.199 and 1.200.)
In the various orders of the Deciduates, the placenta differs considerably both in outer form and internal structure. The extensive investigations of the last ten years have shown that there is more variation in these respects among the higher mammals than was formerly supposed. The physiological work of this important embryonic organ, the nutrition of the foetus during its long sojourn in the womb, is accomplished in the various groups of the Placentals by very different and sometimes very elaborate structures. They have lately been fully described by Hans Strahl.
The phylogeny of the placenta has become more intelligible from the fact that we have found a number of transitional forms of it. Some of the Marsupials (Perameles) have the beginning of a placenta. In some of the Lemurs (Tarsius) a discoid placenta with decidua is developed.
While these important results of comparative embryology have been throwing further light on the close blood-relationship of man and the anthropoid apes in the last few years (Chapter 1.15), the great advance of paleontology has at the same time been affording us a deeper insight into the stem-history of the Placental group. In the seventh chapter of my Systematic Phylogeny of the Vertebrates I advanced the hypothesis that the Placentals form a single stem with many branches, which has been evolved from an older group of the Marsupials (Prodidelphia). The four great legions of the Placentals—Rodents, Ungulates, Carnassia, and Primates—are sharply separated to-day by important features of organisation. But if we consider their extinct ancestors of the Tertiary period, the differences gradually disappear, the deeper we go in the Cenozoic deposits; in the end we find that they vanish altogether. The primitive stem-forms of the Rodents (Esthonychida), the Ungulates (Chondylarthra), the Carnassia (Ictopsida), and the Primates (Lemuravida) are so closely related at the beginning of the Tertiary period that we might group them together as different families of one order, the Proplacentals (Mallotheria or Prochoriata).
Hence the great majority of the Placentals have no direct and close relationship to man, but only the legion of the Primates. This is now generally divided into three orders—the half-apes (Prosimiae), apes (Simiae), and man (Anthropi). The lemurs or half-apes are the stem-group, descending from the older Mallotheria of the Cretaceous period. From them the apes were evolved in the Tertiary period, and man was formed from these towards its close.